Flora of New Zealand Ferns and Lycophytes
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FLORA OF NEW ZEALAND FERNS AND LYCOPHYTES SALVINIACEAE P.J. BROWNSEY & L.R. PERRIE Fascicle 10 – MARCH 2015 © Landcare Research New Zealand Limited 2015. Unless indicated otherwise for specific items, this copyright work is licensed under the Creative Commons Attribution 3.0 New Zealand license. Attribution if redistributing to the public without adaptation: “Source: Landcare Research" Attribution if making an adaptation or derivative work: “Sourced from Landcare Research" See Image Information for copyright and licence details for images. CATALOGUING IN PUBLICATION Brownsey, P.J. (Patrick John), 1948- Flora of New Zealand [electronic resource] : ferns and lycophytes. Fascicle 10, Salviniaceae / P.J. Brownsey and L.R. Perrie. -- Lincoln, N.Z. : Manaaki Whenua Press, 2015. 1 online resource ISBN 978-0-478-34779-1 (pdf) ISBN 978-0-478-34761-6 (set) 1.Ferns -- New Zealand - Identification. I. Perrie, L.R. (Leon Richard). II. Title. III. Manaaki Whenua- Landcare Research New Zealand Ltd. UDC 582.394.771(931)DC 587.30993 DOI: 10.7931/B1H59T This work should be cited as: Brownsey, P.J. & Perrie, L.R. 2015: Salviniaceae. In: Breitwieser, I.; Heenan, P.B.; Wilton, A.D. Flora of New Zealand — Ferns and Lycophytes. Fascicle 10. Manaaki Whenua Press, Lincoln. http://dx.doi.org/10.7931/B1H59T Cover image: Azolla rubra, plants floating on the surface of standing water with irregularly branched stems bearing densely imbricate bilobed floating leaves Contents Introduction..............................................................................................................................................1 Taxa Salviniaceae Martinov ....................................................................................................................... 2 Azolla Lam. ........................................................................................................................................2 Azolla pinnata R.Br. ........................................................................................................................... 3 Azolla rubra R.Br. .............................................................................................................................. 5 Salvinia Ség. ......................................................................................................................................6 Salvinia ×molesta D.S.Mitch. .............................................................................................................7 References ........................................................................................................................................... 10 Acknowledgements ...............................................................................................................................12 Maps .....................................................................................................................................................13 Index .....................................................................................................................................................15 Image Information .................................................................................................................................16 Introduction The family Salviniaceae is represented in New Zealand by two genera, Azolla and Salvinia. Azolla has one indigenous and one naturalised species, whilst Salvinia is represented by a single species which has been declared a noxious weed. Azolla rubra is indigenous, occurring in slow-moving streams and ponds throughout the North Island and the eastern half of the South Island. Azolla pinnata was first recorded in 1969, and is largely confined to the northern North Island, but is now spreading southwards, and has replaced A. rubra in some parts of Northland. Salvinia ×molesta is an extremely aggressive weed of lakes and ponds in Northland and Auckland. Both genera are free-floating, aquatic ferns with highly modified leaves adapted for flotation; they are characteristically heterosporous, producing mega- and microsporangia in hardened sporocarps. Azolla has small leaves and forms red carpets of growth on the water surface. Salvinia has dimorphic leaves arranged in whorls of three, and in S. ×molesta, characteristic egg-whisk-shaped hairs on the floating leaves for buoyancy. 1 Salviniaceae Martinov, Tekhno-Bot. Slovar, 559 (1820) = Azollaceae Wettst., Handb. Syst. Bot. 2, 77 (1903) Type taxon: Salvinia Ség. Aquatic ferns, free-floating. Stems slender, branching, glabrous or hairy; roots either long and pendent, or absent. Leaves either monomorphic and bilobed, with the upper lobe aerial and green and the lower colourless and floating, or dimorphic and arranged in whorls of 3, with two floating and one submerged; not articulated, herbaceous, papillate or with very specialised hairs, non-circinate when young. Veins anastomosing, or absent. Sporocarps globose or conical, sessile or stalked, glabrous or hairy, either attached to the lower lobes of bilobed leaves, or to the submerged leaves, splitting open irregularly, each sporocarp bearing either mega- or microsporangiate sori. Annulus absent. Heterosporous, spores trilete, lacking chlorophyll; one megaspore in each megasporangium and 32–64 microspores in each microsporangium. Megaspores spheroidal, plain to tuberculate; microspores spheroidal, and plain to rugulate. Taxonomy: A family of two genera and about 17 species. The Salviniaceae comprises floating aquatic ferns with highly modified leaves adapted for flotation, mega- and micro-sporangia borne in hardened sporocarps that give rise to mega- and micro-spores, and spores which germinate into greatly reduced gametophytes still attached to the parent spore. Pryer (1999) and Pryer et al. (2004) showed that the heterosporous ferns, Salviniales, are one of three major lineages of “core leptosporangiate” ferns, along with tree ferns and polypod ferns. Nagalingum et al. (2008) subsequently confirmed that Salviniales is divided into two strongly supported families, the Marsileaceae and Salviniaceae, and that within Salviniaceae there is strong support for recognition of two genera, Azolla and Salvinia. Schneider & Pryer (2002) also demonstrated that the spores of Marsileaceae are markedly different to those of Salviniaceae and all homosporous ferns. The morphology of the reproductive structures in heterosporous water ferns has been critically examined by Nagalingum et al. (2006). Some authors (Schneller 1990a; Chinnock 1998; Jones 1998) recognise two separate families, Azollaceae and Salviniaceae, on the basis of the significant differences in the morphology of the two groups, but we follow Smith et al. (2006) and Christenhusz et al. (2011) in recognising just one family. It is represented in New Zealand by one indigenous genus (Azolla) and one fully naturalised genus (Salvinia). Distribution: Widespread in tropical and temperate regions. One native and one naturalised genus with three species in New Zealand; none endemic. Biostatus: Indigenous (Non-endemic). Table 1: Number of species and named hybrids in New Zealand within Salviniaceae Martinov Category Number Indigenous (Non-endemic) 1 Exotic: Fully Naturalised 2 Total 3 Azolla Lam., Encycl. 1, 343 (1783) Type taxon: Azolla filiculoides Lam. Etymology: From the Greek azo (to dry), and ollyo (to kill), a reference to the fact that plants are killed by drying out. Aquatic ferns, free-floating. Stems short, branching, glabrous or sometimes hairy, bearing pendent roots. Leaves monomorphic, bilobed; the upper lobe aerial, green, papillate, containing the cyanobacterium Anabaena in a large cavity on the lower surface; the lower lobe colourless and floating; both lobes glabrous. Veins absent. Sporocarps usually absent in NZ; when present, borne on the first leaf of a lateral branch, in the axil of the lower lobe and protected by the upper lobe; megasporocarps oblong-conical; microsporocarps ovoid to globular, usually larger than megasporocarps. Megaspores solitary in each megasporangium, bearing hair-like filaments at the base and floats (massulae) at the apex. Microspores 32–64 in each microsporangium, mixed with massulae that bear hooked processes (glochidia). Megaspores spheroidal, coarsely tuberculate, perforate; microspores spheroidal, plain. 2 1 Plants regularly 2-3-pinnate; roots with fine lateral rootlets .................... .................... pinnata Plants irregularly branched; roots simple ........................................................................ rubra Distribution: Seven species widespread in tropical and temperate regions (Reid et al. 2006); four occur in North and South America, two in Africa and Asia, and two in Australia, one of which is also native to New Zealand and the other introduced. One naturalised and one native species in New Zealand. Biostatus: Indigenous (Non-endemic). Table 2: Number of species in New Zealand within Azolla Lam. Category Number Indigenous (Non-endemic) 1 Exotic: Fully Naturalised 1 Total 2 Cytology: The base chromosome number in Azolla is x = 22 (Schneller 1990a). Notes: Plants of Azolla are free-floating and capable of covering large areas of slow-moving or still water. They are easily recognised as a red carpet of growth on the water surface. In New Zealand, fertile plants are rarely found, and reproduction is usually vegetative by fragmentation of the lower branches of the plants. Only five collections with sporocarps have been detected in New Zealand herbaria (AK 284894, CHR 168181, 214754, WELT